Seasonal patterns of arthropods occurring on sheltered and

unsheltered pig carcasses in Buenos Aires Province (Argentina)
N. Centeno
, M. Maldonado
, A. Oliva
Centro de Estudios e Investigaciones±Programa de Investigaciones en Interacciones Biologicas (CEI±PIIB),
Universidad Nacional de Quilmes, Roque SaÂenz PenÄa 180, Bernal, B1876BXD Buenos Aires, Argentina
Consejo Nacional de Investigaciones Cienti®cas y Tecnologicas (Conicet)ÐChief of the Laboratorio de Entomologia forense,
Museo argentino de Ciencias naturales, Av. A. Gallardo 470, C1405DJR Buenos Aires, Argentina
Received 8 February 2001; received in revised form 7 February 2002; accepted 8 February 2002
Differences in the succession of insects and other Arthropoda (invertebrate animals with jointed legs), on domestic pig
carcasses placed under a roof and under the open sky have been studied in Buenos Aires Province, Argentina (latitude 34845
in all the seasons of the year. Faunal associations proved different for each treatment in winter: the common bluebottle
Calliphora vicina was found in both, but on the sheltered carcass Cochliomyia macellaria and the rare Phaenicia cluvia were
found as well. In the fall, the difference between sheltered and unsheltered carcasses was small (six species on the former and
®ve species on the latter); in spring and summer, the difference was negligible. #2002 Elsevier Science Ireland Ltd. All rights
Keywords: Cadaveric succession; Sheltered corpses; Calliphoridae; Neotropical fauna; Forensic entomology
1. Introduction
Decay of a dead body is greatly in¯uenced by organisms
which feed upon the body in the different stages of decom-
position. The knowledge of this succession is an important
tool in forensic studies to estimate the interval since death
from the species of organisms found on the body. The most
frequent and numerous living beings to be found on a dead
body are the joint-legged invertebrates comprised in the
phylum Arthropoda, and among these, the class Insecta.
Among the many large groups (orders) of insects, the ¯ies
(Diptera) and the beetles (Coleoptera) have been given the
greatest attention as being instrumental in recycling animal
remains [1]. Our survey is based mainly on insects.
Forensic entomology is based on the knowledge of asso-
ciation of certain species with a given stage of decay, and of
the life-cycles of these species. Insects appear on corpses at
different stages of their development, and in species of the
greatest importance for forensics, such as blow¯ies, the
different life stages may have quite different anatomical
structures. An adult ¯y lays eggs; from these a worm-like
larva (maggot) will hatch; the larva feeds voraciously,
increasing its size at a remarkable rate; when feeding is
completed, the larva becomes sluggish while its gut contents
turn into reserve fat [2]; then the larva will bury itself in the
soil or otherwise seek to hide (a few species may remain on
the feeding substrate at this point of development) and turn
into an immobile pupa, inside a case (puparium); the pupa
turns into a ¯y that must emerge from the puparium, and
often from the soil, before spreading its wings [3].
Insects have outer skeletons made up of plates secreted by
their dermis (cuticle). As cuticle stretches only to a moderate
degree, a growing larva has to discard it, a process called
moult. Breathing is performed through a pair of spiracles in
the rear end of the body and a pair of anterior spiracles that
appears after the ®rst moult. Spiracles are useful to deter-
mine the larval instar, of which there are three: ``larva I''
between hatching and the ®rst moult, ``larva II'' between
larval moults, ``larva III'' between the last larval moult and
pupation. The puparium is derived from the cuticle of the
Forensic Science International 126 (2002) 63±70
Corresponding author. Tel.: ‡54-11-4365-7100x225;
fax: ‡54-11-4365-7182.
E-mail address: (N. Centeno).
0379-0738/02/$ ± see front matter # 2002 Elsevier Science Ireland Ltd. All rights reserved.
PII: S0 3 7 9 - 0 7 3 8 ( 0 2 ) 0 0 0 3 7 - 3
larva III, which hardens into a barrel-shaped brown colored
Insects and other arthropods colonize the body from the
®rst stages of decay, in successive waves; each wave modi-
®es its own substratum, which has the effect of making it
attractive to the next one [4].
Payne [1] studied communities of arthropods associated
with stages of decomposition on the domestic pig Sus scrofa
L.; he exposed carcasses to arthropod activity and studied
decay stages and faunal colonization patterns with the
passing of time. Studies of arthropod succession were
performed in several parts of the world [5±10]. There is a
precedent in [4] for experiment with a roof covering the
corpse to keep off the rain, but the results were not contrasted
with those from an unsheltered one. Also, the animal model
used in [4] was guinea pig, much smaller in size than any
human corpse. In forensic experiments, pig carcass is widely
used because the animal model must closely approximate the
pattern of human decomposition [11]. Pigs are easy to
obtain, their digestive tract resembles the human one (both
species being omnivorous), and the fact that it is often used
as a model makes easier and more accurate any comparing of
results. More about experimental research on forensic ento-
mology in [11]; the history of forensic entomology is
reviewed in [12].
In Southern America, research is being done in Brazil;
results from rat carcasses [13] and from pig carcasses [8±10]
are available. In Argentina, some descriptive reports on
cadaveric insects has been published [8], but up to now,
no attempt was made to determine faunal associations
related to stages of decomposition. Understanding of these
associations is useful, as they contribute to estimation of
postmortem interval and other cases, like children neglected
by their parents, corpse movement after death, etc. [14,15].
The following paper compares decay stages and cadaveric
fauna on sheltered and unsheltered pig carcasses, for each
season of the year. Domestic pig S. scrofa was selected as an
animal model [11].
The area where the experiment was performed has a
warm-temperate, humid climate. Usually, summer (Decem-
ber±March) is warm, winter (June±September) mildly cold
with occasional frosts, fall is subject to strong southeastern
winds with cold and drizzle, which may last for several days;
falls of rain are frequent in spring.
2. Material and methods
The experiments were performed on a ®eld (property of
the Universidad Nacional de La Plata), about 30 ha in sur-
face, located at Santa Catalina (34845
S, 58825
W), some
20 km SE from the city of Buenos Aires, Argentina. The
®eld was sown with maize; alongside there is a wood of
elmtrees and tala trees (a local species of Celtis). The area of
work was 500 mlong Â20 mwide, set between the ®eld and
the wood; it had a lowvegetation of thistles, nettles and a few
shrubs. Logs, fallen branches, leaf mould and small amounts
of inorganic garbage were found in the area.
The climate in this part of the country is humid warm-
temperate. Summers are usually warm, varying from dry
sunny days when the maximum temperatures approach
40 8C, to sudden showers of rain which may lower tem-
peratures to some 18 8C. Sultry weather may prevail for
several days running. Minimum temperatures keep high
even at night, rarely descending below 18 8C except after
a fall of rain. High minimum temperatures make for an
extremely small diel variance, sometimes hardly more than
1 8C. Winters are mildly cold, with frosty spells which are
usually short and interspersed with thaws in which max-
imum temperature may attain 17 and 18 8C. Southeastern
winds, sustained for several days, causing rain and drizzle,
occur at any time of the year, but with greatest frequency in
the fall. Spring and fall very unstable; both frosts and spells
of sultry warmth may occur, interspersed with cool to mildly
warm weather. Annual rainfall for this part of the province is
estimated around 900±950 mm.
For each experiment two wooden cages, 120 cmÂ
80 cmÂ60 cm, were used, one with a wooden roof and
the other one roo¯ess. The cages had a wire mesh 2.5 cm
wide. Inside each cage the body of a domestic pig weighing
15±17 kg, was placed. The pigs were killed by a stab in the
heart (the usual procedure of commercial butchers) 1 h
before exposure, and kept in a plastic bag to avoid insect
infestation before the setting of the experiment. Around each
cage and at a distance of 40 cm were placed six pitfall traps,
two along each of the longer sides, one on each of the shorter
ones. Pitfall traps consist of a small pit dug in the soil,
containing a vial without a bait. For control, six further
pitfall traps were placed at a distance of some 4 m, following
the same spatial pattern. The cages were separated by a
distance of approximately 4 m.
The stages of decay were de®ned as in [1], with slight
modi®cation, as follows:
1. Fresh stage: no decomposition odor or swelling.
2. Bloated stage: abdomen bloated; bubbles of blood at the
nose and anus; in later part of this stage odor becomes
noticeable and ¯uids seep out. It clearly ends when the
corpse de¯ates.
3. Active decay stage: odor strong; liquefaction and
disintegration noticeable; usually skin pierced by
feeding insects; ¯esh may have been removed from
the ®rst points of attack, such as mouth and eyes, in
extreme cases from most of the head.
4. Advanced decay stage: odor begins to fade; most of the
¯esh has disappeared, some soft tissue still found in the
5. Remains stage: odorous disappear; only bones, hair and
remnants of dried skin remain.
The carcasses were visited daily at the beginning of the
experiment (fresh stage), and afterwards at varying inter-
vals, from 3±4 days, according to season (Figs. 1±4).
64 N. Centeno et al. / Forensic Science International 126 (2002) 63±70
Fig. 1. Successional pattern in fall, for forensically important insects; Calliphoridae (blow¯ies), Muscidae (house¯y and relatives), Fannidae
(latrine ¯ies), Silphidae (carrion beetles), Staphylinidae (rove beetles), Histeridae (hister beetles), Dermestidae (skin beetles) and Cleridae
(ham beetles).
Fig. 2. Successional pattern in winter, for forensically important insects; Calliphoridae (blow¯ies), Muscidae (house¯y and relatives),
Fannidae (latrine ¯ies), Silphidae (carrion beetles), Staphylinidae (rove beetles), Histeridae (hister beetles), Dermestidae (skin beetles) and
Cleridae (ham beetles).
N. Centeno et al. / Forensic Science International 126 (2002) 63±70 65
On each visit, samples of arthropods on the body, under it, in
¯ight and in the pitfalls were taken. Catching devices
included forceps, spoons, entomological nets, aspirators
and other implements. In advanced stages of decomposi-
tion, samples of soil were taken, both from under the body
and around it.
Ambient and carcass temperatures were measured, the
latter by means of a 15 cm probe thermometer fully inserted
in the anus. A t-test [16] was performed comparing both
sheltered and unsheltered mean body temperatures, from the
bloated to the beginning of advanced decay stages (which
were the maximummaggot activity periods) for each season.
Fig. 3. Successional pattern in spring, for forensically important insects; Calliphoridae (blow¯ies), Muscidae (house¯y and relatives),
Fannidae (latrine ¯ies), Silphidae (carrion beetles), Staphylinidae (rove beetles), Histeridae (hister beetles), Dermestidae (skin beetles) and
Cleridae (ham beetles).
Fig. 4. Successional pattern in summer, for most forensically important insects; Calliphoridae (blow¯ies), Muscidae (house¯y and relatives),
Fannidae (latrine ¯ies), Silphidae (carrion beetles), Staphylinidae (rove beetles), Histeridae (hister beetles), Dermestidae (skin beetles) and
Cleridae (ham beetles).
66 N. Centeno et al. / Forensic Science International 126 (2002) 63±70
Fresh and remains stages were not considered for analysis
because carcass temperature did not rise above air tempera-
ture, as observed in [1].
Blow¯y eggs and half of collected maggots were reserved
for lab rearing; the remaining larvae were killed in hot water
at 80 8C and preserved in ethanol 70%. Adult ¯ies were
killed with ethyl acetate vapor and placed in paper envel-
opes; the remaining arthropods were kept in 70% ethanol.
Cultures were kept in plastic jars 7 cm in diameter; beef was
used as food, placed on pieces of aluminiumfoil, on a base of
perlite as suggested in [17]. The cultures were placed in a
hothouse at 24 8C until completion of development, to
con®rm determination through the adults. Determinations
were based on [18±20].
This experiment was repeated in the four seasons of the
year beginning in the winter (June±September) of 1998 and
ending in the summer (December±March) of 2000.
Since there was a single set of data for each combination
of variables, no statistical analysis of the results concerning
insect sampling was feasible. The results of analysis of
internal temperature measurements and their correlation
with external conditions are shown in Table 3.
3. Results
The ®rst blow¯ies eggs were always found during the
fresh stage for both treatments: after the ®rst day in the fall,
after the second day in winter, within the ®rst day in spring
and summer. In later stages, fresh egg-laying by blow¯ies
occurred (recolonization): in fall, this was observed until day
41 on the sheltered and day 48 on the unsheltered carcass; in
winter, until day 15 on the sheltered and day 45 on the
unsheltered carcass; in spring and summer, there was reco-
lonization until day 5 for both treatments.
The ®rst records of larvae III were made on the third day
after the carcass was exposed, for both treatments, in spring
and in summer. In fall, the ®rst record was made on the
eighth day, for both treatments. Only in winter a difference
between treatments showed itself; the ®rst record of larvae
III on the sheltered carcass was in the eighth day, on the
unsheltered carcass, on the day 14.
Figs. 1±4 show the faunal associations, how each relates
to decomposition stages and to day accumulation from the
day of exposition onwards, for each season and for each
treatment. The largest segment of the association was made
Table 1
Insects of forensic importance collected on corpses
Diptera (true flies) Calliphoridae (blowflies) C. vicina Robineau-Desvoidy
C. nigribasis Macquart
P. sericata (Meigen)
P. cluvia (Walker)
Paralucilia pseudolyrcea (Mello)
Chrysomya albiceps (Wiedemann)
C. chloropyga (Wiedemann)
C. megacephala F.
C. macellaria F.
Sarconesia chlorogaster (Wiedemann)
Muscidae (housefly and its relatives) Ophyra argentina Bigot
Morellia spp.
Muscina stabulans(Fallen)
M. asimilis Fallen
Musca domestica L.
Fannidae (latrine flies) Fannia fusconotata (Rondani)
Phoridae (scuttle flies) Megaselia scalaris (Loew)
Spiniphora bergenstammi (Mik)
Puliciphora rufipes Silva Figueroa
Coleoptera (beetles) Silphidae (carrion beetles) Hyponecrodes sp.
Cleridae (ham beetles) Necrobia ruficollis F.
N. rufipes De Geer
Histeridae (hister beetles) Saprinus patagonicus (Blanchard)
Hister sp.
Staphylinidae (rove beetles) Creophilus maxillosus L.
Paederus sp.
Dermestidae (skin beetles) Dermestes ater De Geer
D. maculatus De Geer
Hymenoptera Formicidae (fire ants) Solenopsis sp.
N. Centeno et al. / Forensic Science International 126 (2002) 63±70 67
up of blow¯ies (Calliphoridae), although ¯ies of the house¯y
family (Muscidae) and latrine ¯ies or lesser house¯ies
(Fanniidae) also occurred.
The fresh stage was characterized by blow¯ies eggs and
larvae I; the blow¯ies larvae II and III was the main
components in the bloated stage together with carrion
beetles (Silphidae) adults; active decay stage showed mostly
larvae III of blow¯ies and the coexistence of carrion beetle
larvae and adults; advanced decay stage showed a majority
of blow¯y postfeeding larvae leaving the body and carrion
beetle larvae (no adults); in the remain stage, beetles were
the most abundant group, although no family appeared to be
especially associated to this stage.
The families of Diptera and Coleoptera that were col-
lected on the carcasses are shown in Table 1. Taxa, that were
present in the control traps but not in the carcass are listed in
Table 2. For the analysis of the Dipteran faunal associations,
larvae alone were considered. Adults ¯ying nearby were not
taken into account. Insects trapped in control traps were not
taken into account, save for rove and carrion beetles which
have been recorded as associated with corpses [1,4].
In the fall, slight differences were observed between
sheltered and unsheltered carcasses. On the later, Calli-
phora vicina, C. nigribasis, Phaenicia sericata, P. cluvia
and Cochliomyia macellaria occurred; on sheltered carcass,
the same precedent ®ve species occurred, and C. albiceps
which is usually associated with warmweather was found as
well. In winter, the difference was more marked; on the
unsheltered carcass only C. vicina (the most common
species at this time of the year) occurred, while on the
sheltered carcass C. vicina, P. cluvia and C. macellaria were
In the spring, no difference between sheltered and unshel-
tered carcasses was observed. In both cases, C. macellaria,
C. albiceps, P. sericata, P. pseudolyrcea and C. chloropyga
were found. In the summer, there was a small number of
species, without signi®cant differences between sheltered
and unsheltered carcasses; decomposition took a short space
of time (22 days), C. albiceps and C. macellaria occurred,
and in smaller numbers C. megacephala. From the ®rst day,
Table 2
Groups present only in control traps without forensic importance
Coleoptera (beetles) Carabidae, Scarabaeidae, Coprinae,
Lampyridae, Tenebrionidae
Heteroptera (true bugs) Reduviidae, Nabiidae, Pyrrhocoridae
Homoptera (leaf hoppers) Fulgoroidea, Cicadelidae, Aphidoidea
(ants and bees)
Attinae (leaf-cutting ants),
Apidae (bees)
Dictyoptera (cockroaches) Blattodea (cockroaches)
Lepidoptera (butterflies) Larvae (caterpillar)
Collembolla (springtails) Isotomidae, Onychiuridae, Poduridae
Chilopoda Centipedes
Diplopoda Millipedes
Opiliones (harvestmen) Pallpatores and Laniatores
Araneae (spiders) Lycosidae (wolf spider)
Crustacea Isopoda (pill bugs)
Mollusca Gasteropoda (snails)
Fig. 5. Winter temperatures. Comparison of sheltered and unsheltered corpse temperatures. Interval between broken lines showing interval of
days analyzed with t-test of difference between both mean temperatures. Signi®cance level, a ˆ 0:05; S: signi®cant at the 0.05 level.
68 N. Centeno et al. / Forensic Science International 126 (2002) 63±70
C. albiceps was the most numerous, although the ®nding of a
single larva I of C. macellaria on the day after exposition
suggests that this species may be the ®rst to colonize the
Comparison between sheltered and unsheltered mean
carrion temperatures, on bloated and active decay stages,
showed signi®cant differences only in winter (P < 0:05; 18
d.f.; see Fig. 5). In contrast, results of the test (Table 3)
showed no signi®cant differences for the fall, spring or
4. Discussion
Stages of decomposition: Differences between sheltered
and unsheltered carcasses were clear-cut. This might be
attributed to the effect of the roof. The trend in fall and
in winter is towards a shorter bloated stage in sheltered
carcasses. Both active decay and advanced decay were
shorter for unsheltered carcasses in winter, but this might
be due to an overlap of the beginning of active decay with the
end of the bloated stage; in the unsheltered experiment, the
bloated stage remained well-de®ned until the ®rst postfeed-
ing larvae appeared (postfeeding larvae usually appear from
the advanced decay stage on). These differences may be
explained by the fact that the roof protects the carcass from
rain and frost. This idea is also supported by the higher
temperatures recorded for the sheltered corpse (Fig. 5); this
may possibly accelerate decomposition.
In spring, on the other hand, active decay and advanced
decay were shorter in the unsheltered carcass than in the
sheltered one. This may be due to the reduction in the
insolation caused by the roof, making for a longer decay
period, although records of temperatures do not show lower
values (see Table 3).
At a lower latitude, as in Brazil [13], the stages of
decomposition were longer in winter than in any other
season. In Buenos Aires Province, however, the difference
was more clear-cut, e.g. the active decay stage lasted 3 days
in summer and 19 days in the fall. Similar differences may be
remarked for the other stages (Figs. 1±4).
The ®rst stages showed important differences both for
each season and between treatments. In the fall, the fresh
stage was shorter than in winter, but comparable for both
carcasses, and distinctly longer than in spring or summer.
The period between the bloated stage and the end of active
decay stage, both in the fall and in winter, was comparable
but not equal for both treatments, being shorter for the
sheltered carcass. In the fall, the advanced decay stage
was distinctly longer for the unsheltered carcass, while in
winter values were similar for both treatments.
In spring, succession was very quick, in 22 days the
carcasses had arrived at the remains stage [1]. This might
be an effect of insolation which increases temperature. Afact
which supports this supposition is that the higher tempera-
ture values were recorded for the unsheltered carcass; on the
following day after beginning the experiment, there was a
temperature difference of 6 8C between the two treatments.
This acceleration in the successional rate affected all the
stages, the unsheltered carcass arrived at the remains stage 5
days before the sheltered one. In summer, no signi®cant
differences were recorded between the two treatments; as in
spring, the remains stage was attained in 22 days.
The differences found in the assemblage of Calliphoridae
in the cold months (April±August), have been considered as
the effect of the roof on the process of decomposition. On the
sheltered corpses, the presence of summer blow¯ies species
[21] such as C. albiceps, P. cluvia and C. macellaria, as well
as the fast development of the larvae III (in comparison with
other treatment), could be due to the protection of the roof.
On the other hand, in the unsheltered carcass slow decom-
position allowed recolonization by blow¯ies until a later
date than on the sheltered one.
The families of Coleoptera (beetles) showed different
seasonal patterns. Skin beetles (Dermestidae) were found
from summer to the beginning of the fall. Carrion beetles
(Silphidae) occur along the whole year, except in summer.
Rove beetles (Staphylinidae) showed no seasonal differ-
ences. However, in the fall they appeared earlier on the
sheltered carcass; this fact may be due to a microclimate
generated by the roof.
In conclusion, differences between sheltered and unshel-
tered carcasses were found to be signi®cant for the succes-
sion of necrophagous insects. A shelter or cover affects the
decomposition process, making it shortest in cold seasons
(fall/winter) and longest in warm seasons (spring/summer),
as compared with an exposed condition of a corpse. The
successional pattern and the species recorded, show differ-
Table 3
Results of the t-test to compare mean corpse temperatures between both sheltered and unsheltered pig carcass
Season Day interval U mean (8C) S mean (8C) d.f. Significance
Fall 5±43 18.73 18.79 22 NS
Winter 8±43 20.50 23.80 18 S
Spring 1±10 31.87 29.50 7 NS
Summer 1±7 41.75 40.75 6 NS
U mean: mean corpse temperature (8C) for unsheltered pig carcass; S mean: mean corpse temperature (8C) for sheltered pig carcass; d.f.:
degree of freedom of the test; signi®cance level, a ˆ 0:05, S: signi®cant at the 0.05 level (P < 0:05) and NS: not signi®cant at the 0.05 level
(P > 0:05).
N. Centeno et al. / Forensic Science International 126 (2002) 63±70 69
ences due to sheltering during cold seasons. This is of
interest, because in criminal cases there may be attempts
to conceal the body with branches or some other covering.
The authors would like to thank to Dr. Carlos Naranjo,
Director of the Instituto Fitotecnico de Santa Catalina, and
Enrique Michel, foreman of the same Institute; Dr. Arturo
Roig, from the Museo Argentino de Ciencias Naturales, for
determination of Hymenoptera, and Bruno Frasanito, from
the Universidad Nacional de Quilmes, for their help in the
®eld studies. This work was made with the ®nancial support
from the Universidad Nacional de Quilmes.
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